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APES Evolution Lab Name_________________________

What Does a T. Rex Taste like? and Island Biogeography and Evolution

Go to and read and follow the ALL directions.

Features Table: As you explore Folder 4, fill in the data tables below, using a +, -, or ?.

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[pic] example cladogram or phylogenic tree

So what does a T. Rex taste like?

Island Biogeography and Evolution:

Solving a Phylogenetic Puzzle With Molecular Genetics

Introduction

Ever since Charles Darwin formulated his hypothesis on how the finches of the Galapagos Islands evolved into 13 species, islands have been a prime target for the study of evolution. By their very nature, islands are isolated and are essentially a living laboratory of evolution. In this investigation you should be familiar with the terms speciation, geographic isolation, gene flow, gene pool, and reproductive isolation. You will work with real data from real populations. The data will include observations of lizard morphology (body form), geological age estimates of various islands in the Canary Island Archipelago, geographic distances, and genetic distances based on nucleotide base differences in DNA between different populations of lizards.

Background

There are many populations of lizards living on the Canary Islands, and photographs of various examples can be found in Thorpe (1989). The Canary Islands form an archipelago of seven volcanic islands just west of the African continent (Map 1). The island chain starts about 85 km (50 miles) from the continent following a fault line of the Atlas Mountain in northern Africa. Geologists theorize that a geologic hot spot of upwelling magma has been drifting westward for the past 20 million years gradually forming the islands as it moves. Thus, the furthest east island, Lanzarote is oldest while the smaller western island, Hierro is the youngest, about 0.5 million years old. Volcanic islands are particularly good laboratories for evolutionary science because they can be dated accurately using radioactive isotope decay and because they start out as lifeless masses of rock emerging from the sea.

The development of ecosystems on volcanic islands is somewhat unpredictable. However, ecological succession does occur first with pioneer organisms that gradually alter the environment until a stable climax community is established. What is unpredictable is: What plant and animal species will colonize these new environments? Much of this is left to climate, proximity to other land masses, and of course, chance. This investigation deals with three species of lizards of the genus Gallotia, and within one of these species, Gallotia galloti, four separate island populations. The arrival of the Gallotia lizards was probably by rafting (See Map 1), that is, rafts of natural vegetation are often washed out to sea when high river levels cause river banks to collapse, carrying away both plants and clinging animals alike. Colonization by airborne organisms such as insects and birds usually occurred during storms. In any case, there are some general principles of island colonization which follow:

1) The closer the island is to another land mass, the higher the probability of colonization.

2) The older the island, the more likely it will be colonized.

3) The larger the island, the more species are likely to be established.

4) The geographic isolation reduces gene flow between populations.

5) Over time, colonial populations become genetically divergent from their parent population due to natural selection, mutation, and/or genetic drift.

Map 1. The Canary Islands Archipelago. (Redrawn from Anguita, 1986).

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Problem: Evolution biologists have been faced with an interesting problem. What is the phylogenetic history of the three species and seven populations of Gallotia lizards on the islands? Does the presence of four morphologically different populations of G. galloti on the four western most islands (Map 2) imply continuing evolution? In this investigation, you will use data from geography, geological history, morphology (body size), and molecular genetics to develop answers to these questions.

Map 2. On the map below, three species of lizards are shown, Gallotia atlantica, G. stehlini, and G. galloti. G. galloti has colonized the four western most islands and each population is morphologically distinct from the other. (Redrawn from R.S. Thorpe, 1993.)

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Procedure – complete drawings and answer all questions, in complete sentences, for each part of this lab activity. You may type your responses and draw your cladograms (phylogenic trees) using MS Word or by hand, your choice. It must be turned in by the end of the period.

Part I: Phylogeny Based On Geographic Distance

Using Map 1, measure the distances in kilometers of each island to the mainland (Africa). List these distances on a separate page. Include the following islands: Lanzarote, Fuerteventura, Gran Canaria, Tenerife, Gomera Palma, and Hierro.

1. Which island would have been colonized first and which last? Explain why you think so.

2. Using Map 2 and your geographic reasoning, diagram on a separate page a hypothetical phylogenetic (family) tree of the three species and three additional populations of G. galloti. Use what you learned from What does a…. Label your end branches with the following population names:

|atlantica |stehlini |galloti |galloti |galloti |galloti |

| | |Tenerife |Palma |Gomera |Hierro |

Part II: Phylogeny Based on Geological History

Check your hypothetical phylogenetic tree against the geological data in the Table 1. The maximum age of each island was estimated by sampling volcanic rocks found on all islands. The ratio of radioactive potassium to its breakdown product argon was used to estimate the age of the rocks. Redraw you phylogenic tree using this additional data, if changes are necessary.

Table 1. Maximum age in millions of years for the Canary Islands (from F. Anguita, 1986).

|Lanzarote & |Gran Canaria | | | | |

|Fuerteventura | |Tenerife |Gomera |Palma |Hierro |

|24.0 |17.1 |15.1 |5.3 |2.0 |0.8 |

1. Explain how the data in Table 1. supports your first phylogeny diagram. Or, what changes did you make and why?

Part III: Phylogeny Based On Morphology

Study the drawings from each lizard population in Figure 1 and compare and contrast their body size with the distribution on Map 2. To be sure differences are genetic, not ecological, researchers collected individuals from all island populations and bred and raised them in captivity. Their offspring still displayed differences according to their parental characteristics. Draw a new phylogeny chart based on morphological similarities and differences.

1. Compare your two (or three) phylogeny charts. Describe how they are different.

Figure 1. The relative sizes of typical lizards from each population are shown. Lizards on Lanzarote and Fuerteventura are essentially the same. (Redrawn from R.S. Thorpe, 1994.)

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Part IV: Phylogeny Based On Molecular Genetics

Recent studies by R. S. Thorpe (1993, 1994) have attempted to support various phylogenetic hypotheses by comparing genetic differences among the populations of the Gallotia lizards on the Canary Islands. The gene for cytochrome b which is coded by DNA found in every cell's mitochondria was used in this study along with DNA from other genes. Cytochrome b is an important substance for cell metabolism and has probably been around since the first prokaryotes. Changes in its nucleotide base sequence (A, T, C, and G) that do not disrupt the gene's function provide us with a kind of evolutionary clock. The rate of mutational changes due to pairing errors is relatively constant. The chances for such mutations are the same for any of these bases. This means that the more time, the more changes. When two populations are isolated and gene flow between them is restricted, the mutational differences accumulate over time. The longer the isolation the greater the difference.

Thorpe and his colleagues used restriction enzymes to cut the DNA and gel electrophoresis to separate the fragments. Radioisotope tagging eventually led to the sequencing of the samples of DNA for each of the seven populations. Thorpe tested two populations on Tenerife to see if ecological differences were part of the story. He felt that because Tenerife is moist and lush in the north while arid and barren in the south, populations on that island might have some genetic differences. Also, he wondered if Tenerife was supplying colonizing lizards from two different directions. The results for Thorpe's tests appear in Table 2 and Table 3 of this investigation.

Your task is to review the differences between all pairings of the seven populations and use that data to construct a final phylogenetic tree based of genetic similarities and differences.

Procedure for Part IV:

There are twenty-one different pair combinations possible using seven populations. The pairings are listed on Table 3. Look for low numbers which express more genetic similarity and imply more recent common ancestry. Pairs that produce high numbers are said to have greater genetic distance between them. In other words, large numbers imply they are less genetically alike and have more distant ancestry. On a phylogenetic tree, distant ancestry is expressed by low branches while more recently evolved and more recent ancestry are on the higher branches.

Table 2 Cytochrome b DNA bases for the seven populations of Gallotia Lizards

Table 3 Cytochrome b DNA sequence differences for the seven populations of Gallotia Lizards

  |1.

G.stehlini |2.

G. atlantica |3.

G. galloti P. |4.

G. galloti N. T. |5.

G. galloti S.T. |6.

G. galloti G. |7.

G. galloti H | |1. G. stehlini | 

|  |  |  |  |  |  | |2. G. atlantica |36 | 

|  |  |  |  |  | |3. G. galloti Palma |41 |25 |  |  |  |  |  | |4. G. galloti N. Tenerife |40 |21 |8 |  |  |  |  | |5. G. galloti S. Tenerife |40 |19 |10 |6 |  |  |  | |6. G. galloti Gomera |45 |22 |13 |19 |15 |  |  | |7. G. galloti Hiero |49 |28 |17 |21 |15 |4 |  | |Final Interpretations and Conclusions

1. On Table 3, large numbers imply that pairs of populations have more distant ancestry. Why is this?

2. How many base pair differences do you think separates any two species of these lizards? Give an example to support your answer.

3. Which two populations are most closely related? Justify your answer.

4. Why should you expect the populations S. Tenerife (ST) and N. Tenerife (NT) to have fewer differences than other pairings?

5. Which population is least related to the rest? Why do you say so?

6. Draw a phylogeny chart using genetic similarities and differences found in Table 3. Compare it to the phylogeny based on the geologic age of the islands.

7. What difference is there between the two phylogenies?

8. Would you say that G. stehlini is the ancestor of G. atlantica or vice versa or are they ancestors at all? Explain you reasoning.

9. Predict what is likely to happen to the four populations of G. galloti on the four western islands and identify what conditions will support your predictions.

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